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Wednesday, June 10, 2015

101 ancient Eurasian genomes (Allentoft et al. 2015)

It'll take me a while to digest all of the information in this massive new Allentoft et al. paper. But I've already noticed that, just like in Haak et al. 2015, the Yamnaya samples are again from the eastern half of the Yamnaya horizon. This time, however, not all of the Yamnaya individuals carry Y-haplogroup R1b; one of the five samples belongs to Y-haplogroup I2a (see here).

So I'm wondering what more westerly Yamnaya sites will reveal in the future, considering the predominance of Y-haplogroup R1a among the Corded Ware individuals sampled to date, and the close genome-wide relationship between the Yamnaya and Corded Ware?

Abstract: The Bronze Age of Eurasia (around 3000–1000 BC) was a period of major cultural changes. However, there is debate about whether these changes resulted from the circulation of ideas or from human migrations, potentially also facilitating the spread of languages and certain phenotypic traits. We investigated this by using new, improved methods to sequence low-coverage genomes from 101 ancient humans from across Eurasia. We show that the Bronze Age was a highly dynamic period involving large-scale population migrations and replacements, responsible for shaping major parts of present-day demographic structure in both Europe and Asia. Our findings are consistent with the hypothesized spread of Indo-European languages during the Early Bronze Age. We also demonstrate that light skin pigmentation in Europeans was already present at high frequency in the Bronze Age, but not lactose tolerance, indicating a more recent onset of positive selection on lactose tolerance than previously thought.

Another observation, RISE486 which is labeled as one of the Remedello samples, dates to 1773 cal BC. As far as I know the Remedello culture didn't exist anymore at such a late date. According to wikipedia, Remedello ends in 2400 BC. But whatever its exact cultural affiliation, 1773 BC definitely postdates the Bell Beaker period in Northern Italy. And yet this sample was I2, not R1b. Which seems to suggest that Bell Beaker wasn't everywhere dominated by R1b.

"Also, Kurti, it should be safe to consider that these people were recent arrivals to the area, so it doesn't necessarily mean that SW Asian was not present in the area before; rather that these newcomers didn't get it yet."

I doubt it, I really doubt it and I gave a reasoning for why.

Yamna almost completely lacked Atlantic_Med and had very little of Caucasus. Therefore this must have been there native admixture.

And when the majority of their aDNA was already different from that of Yamna than if Red Sea existed in the region obviously they had to have at least few percentage of this Red Sea/SOuthwest Asian.

But they have nothing in all the samples I have yet seen. During Iron Age they appear to get heavy admixture from I would say Iran, still no Red Sea/Southwest Asian.

@Matt

obviouly the Armenian samples are not a good proxy for a Neolithic farmer. My point also was to just show, that this farmer DNA in this regions was more EEF type than nowadays. In all of West Asia Atlantic_Med nowadays does not exceed 15% and the people have more "Southwest Asian" which itself is basically a hybrid of Atlantic_Med with Red Sea. And Atlantic_Med was twice as much in this Bronze Age sample. Thats the point. And it's appearance on the genetic landscape overlaps well with the expansion of Semite speakers.

Regarding J2, it's most certainly from West Asia. J2b does predominate in Iron Age Armenians. The fact that J2a was found in the Iron Age Altai only shows that after Andronovo it wasn't only East Asians who migrated to central Asia, but also West Asians. And therefore I'm not so sure that Tocharian comes from Afanasievo, and not from a West Asian IE population.

Bronze Age/Iron Age is too early for the arrival of the Armenian language.

Also 2500 BC is generally too early for any attested Indo_European language on the region or in general. I am not even sure if it is connected to Indo Europeans. If it is connected to Indo Europeans I have my doubts that the came via the Steppes.

Therefore R1b must have arrived from a different source. Maybe even around Neolithic. I think North Iran is a better quess as you said.

As for David Anthony's theory on the southern admix in Yamnaya and Khvalynsk: Possible. But maybe though this southern admix rather means that archeologists have underestimated the West Asian influence on the steppe. Stanislav Grigoriev has accumulated evidence which in his opinion prove West Asian/Caucasian cultural influence on the steppe already at an early date:

According to him, southern influence is still vague in the 6th millennium BC, at most pointing to a sluggish infiltration of Caucasian populations, without fundamental effect on the local cultural development.

But then, from 5000 BC, he finds the following items pointing to Caucasian influence:polished adzes, maceheads, large blades, tanged arrowheads and flat bottomed pottery.According to Grigoriev, these are ultimately derived from the Hassuna culture. Moreover he also mentions jet beads as Caucasian influence, and the collared ware, which was also found in Ginchi, in the Northeastern Caucasus. And he compares channelled bone plaques from Yarim Tepe (Hassuna level) to similar ones made of boar's teeth in the Northern Caucasus and Mariupol. Lastly, he also compares the arrow straighteners of the steppe to ones from Eastern Anatolia and the Transcaucasus.

Then, after 4500 BC, he finds the new custom of contracted on the back burials, which are also found in the Transcaucasian Eneolithic. Less often also secondary burials, intramural burials and burials without skull, according to Grigoriev all unambigously of southern tradition. Furthermore he mentions as southern cultural influence: the applied decoration on one of the vessels from Khvalynsk, vessels with their sides coloured with ochre, traces of polishing on Sredniy Stog ware, and stone sceptres. He also notes that in the southern steppe polished ware was quite characteristic. Similar to ware from the Transcaucasus are Novodanilovka amphorae with applied knobs, and Grigoriev finds that Novodanilovka and Lower Mikhailovka pottery in general resembles Transcaucasian pottery, and sees similar parallels in the hollow-based arrowheads. He also thinks that the anthropomorphic stelae have early Near Eastern parallels and notes the early appearance of burials under barrows in the Transcaucasus, in Golitsino and Stepanakert. Catacomb burials, which occur on the steppe for the first time at this period, are also found in Northern Mesopotamia and the South Caspian. Lastly he mentions an obsidian dagger from Novodanilovka with eastern Anatolian parallels; the obsidian used stemmed from eastern Anatolia, according to scientific analysis.

Then, as for the Yamnaya period, he thinks that Kemi Oba has strong Caucasus influence. Also he finds Novosvobodnaya metal work in Yamnaya, axes for instance. But he thinks that there's no evidence for the idea that a large West Asian population penetrated into the steppe at the beginning of the Yamnaya culture. Yet he finds polished, Caucasian looking pottery also in the upper levels of Mikhailovka and also stone fortifications, like in the Near East and Maikop. And he regards Usatovo as a mix of steppe, Caucasus and Near Eastern influences on a CT substrate.

Now again back to the conclusions. What we see here is a steady increase of Red Sea like /Southwest Asian genes over time. First from 2% to 4% and now 7% Southwest Asian.And this probably went on until we reach the 14% average nowadays found in the region.

The SSA/Southwest Asian (Red Sea) rises and rises steadly. The only people I see bringing that admixture are Afro_Asiatic speakers because it correspondence well with the appearance of them in the region of Transcaucasus.

What ever You think about Armenian language age ( I think most of these theories are outdated from the beginning of 20th century ) there should be some solid explanation why Yamna and Armenian R1b shows so much similarity. Maybe a common source maybe the Yamna is the parent.

Certainly Sintashta having a North Central European origin is the biggest surprise in this study. The Andronovo samples should look similar, but more mixed with the local populations that were around.

As for the Armenian samples, the "Armenian-like" population that mixed with EHG in the steppe was something more like Tabasaran. In K8 they would have been something around 13/52/32 (WHG/ENF/ANE). Maybe the Bronze Age samples from Armenia will be somehow close to that?

A couple things. Modern Russian area of Yamnaya is not heavily tested in any database FTDNA or elsewhere. Most Russian testers come from NW Russia, areas we know are dense in N1c1 and R1a1. R1b was likely always a minority in this region.

It does kind of look like steppe cultures were running west and south for the hills. I suspect this is due to East Asian tribal incursions from the Altai region.

Alberto, i will like to see the Y-DNA first from other parts of Asia and Sintashta itself! as we don't know the Mutation yet but i suspect it to be R1a-M417, MTdna wise( though i don't know the subclades) Sintashta mtDNA is U2, which is mainly Asian: http://en.wikipedia.org/wiki/Haplogroup_U_(mtDNA)#Haplogroup_U2N1, which was found in Neolithic Europeans and is possibly of Arabian origin: http://en.wikipedia.org/wiki/Haplogroup_N1a_(mtDNA) J1 and J2, also of Near Eastern origin.

But that's not so northern. In relative terms (for the region), yes. But it's still in the expected levels (some 13% WHG, which is what the Armenian-like population from Yamnaya should have. We'll have to see their ENF and ANE levels first, but they could match well the profile needed to form a Yamnaya population with EHG.

David, you know that Sntashta can't be the ancestor of Any Indian Population because of it Autosomal structure and it will not matter if ''Sintashta R1a at the highest resolution will be ancestral to South Asian R1a-Z93.'' because ancient dna from India or other parts Asia will have the R1a in tons before the 2000 BC period! just wait....Alberto,//Nirjhar, but no matter the haplogroup they have, someone scoring some ~70% WHG in K7 must have European origin. There's no way around that.//70% WHG is just astonishing but in case of Autosomes i support what Maju says they are a subject of high change and replacement within few generations so we can't trust them much.

One of the results I am really looking forward to is the analysis of the lone Yamna I2a. Judging by the info from Figure 2 of the Allenhoft paper, this individual had no measurable "farmer type" characteristics (so we can probably rule out Trypilian or Usatovo input). The Verenich take has I2a arising just north of the Alps, and then spreading northward and eastward. This Yamna individual may well have been a descendant of some local HG who entered the PIE-forming process before "farmerization" of most other known I2a's. The Yamna fellow is clearly distinct from the Remedello BA I2a and from the Hungarian I2a's.

It's totally possible I2a somehow made it's way to Russia in the Mesolithic and Neolithic. I2a in Yamnaya is no surprise. Also, it was obviously a small minority.

****A little more than possible now it seems. As for "small minority". That's quite probable, though we still need more ancient DNA from westernmost Yamna to see how small (and of course from adjacent cultural phenomena). And really this doesn't make much difference does it? I'm more interested in the specific identification of the I2a type this individual represents.*****

"That's an interesting and quite controversial theory. But I have to disagree, it's rather not a founder effect. Bell Beakers seem to be uniformly R1b and that's the source of most of modern U106 and P312. CWC in Western Europe was mostly replaced, it's recapitulation is visible just in Frisian/coastal CTS4385/L664, Z283* (Swiss CWC?), Z282*, some Z280(S24902?, Z280*). They are absolutely minor now. One R1b doesn't change anything, look at the patterns and archeology".(Arthur Martyka, answering a post of mine where I quoted Krefter and Davidski from Eurogenes blog, in Human Population Genetics, FB Group).It's time. I won my battle.

I meant north European-like admixed, not in a geographical sense northern. I won't put any money on a concrete origin. But the exciting thing is that, judging from this evidence, R1b in Armenians really came with the Armenian language and from steppe-related people. That is, the R1b-L23 there didn't originate in the South Caspian or in West Asia, and the same holds true for the Armenian language. Conversely R1b-L23 in Yamnaya was from EHG, not from the West Asian element.

This is obviously a heavy blow against the West Asian theory of a PIE Urheimat. It doesn't completely knock it out, but now at most Anatolian and Tocharian might still be from West Asia, at least all the rest is from steppe, and presumably everything is. And if PIE was ultimately from West Asia (which I'd rather not think) it could only be associated with haplogroup J2 (generally speaking, with exceptions of course).

It's fascinating to see how the North European-like admixture initially, in the Bronze Age, was quite substantial (about 25%), then got weaker in the Iron Age (17%) and is very weak nowadays. Apparently they mixed more and more with the locals but were able to preserve their language and ethnicity nonetheless.

It has been observed via physical anthropology that a long headed "Nordic" looking type entered Armenia in the late Bronze Age. Modern Armenian crania differ a lot from this. So maybe this is one of the few times where considerable changes in cranial metrics were paralleled by genetic changes?

I don't see it like that necessarily. As I explained above, the population that mixed with EHG in the Samara region were about as "European" (if you want to use this term, but I'd just say WHG) as these BA Armenian samples. In other words, their level of WHG doesn't prove any steppe or European origin. It actually matches the expectations of the "Caucasus-like" population we're looking for (that should be Lezgin-Tabasaran-Tajik-like).

The fact that they have R1b actually goes against the origin of R1b in EHG. If this "Caucasus-like" population was R1b, then it makes a lot of sense that they brought it to the steppe. Isn't it actually from an older subclade than the one found in Yamnaya?

So it's early to say one way or the other. We need first to see the samples in better detail and then we'll still need more samples, older and from other places. Central Asia proper would be nice to have some answers.

To put their "northerness" in perspective, EEF have from 25-30% WHG up. And it's likely that early farmers from Anatolia had a similar amount. Maybe less, in the 15-20% range. I doubt they would be 0-2% as modern Near Easterners (and Georgians).

That's why the 25% north-German might be quite misleading. It doesn't mean they had any European (including the steppe) admixture. It is just a low amount of WHG (10-15%) which should be expected from that time.

If the samples turn out to be ~25% WHG, then we can start to think about a steppe or European origin.

Whether the old samples from Armenia are closer to North Europe (or even Europe at all) compared to the present day Armenian samples should be easy to check out when Davidski can run stats of the form f4(Mbuti,WHG/North_Europe/Bronze_Age;Armenian,baArm/irArm).

From the f3 and D stats in the paper (some of which I posted above), it doesn't seem very likely that they'll be much closer. The outgroup f3s for Bronze Age Armenian with WHG and SHG and MA1 are close to present day Armenians. A little lower for affinity to WHG and SHG (Bronze Age Armenia further away), and a little higher for Mal'ta, but very close. (baArm f3 for Mal'ta is like Georgians while the f3 for WHG and SHG are like present day Armenians).

Iron Age Armenia has raised WHG / SHG affinity to Bronze Age, reduced Mal'ta, but still WHG / SHG is in the North Caucasus neighbourhood at the highest.

Similar thing if you look at the f3 outgroup stat with Bell Beaker - Armenian (modern) 0.265823 vs Bronze Age Armenian 0.265634. It's not very significant, but the stat is lower, indicating that baArm is further away from Bell Beaker than Armenian is. (irArm has a high stat with Bell Beaker, but that has low SNP overlap, while baArm is close to 300000 SNPs).

Now the highest outgroup f3s for baArm *are* found for North Europeans, like Scots, Icelanders and Lithuanians *but* these are still very low compared with other f3 stats. Tentatively, it seems like the baArm are not very much like anyone, and that the North Europeans (and to a lesser extent other Caucasians) are just the best of a bad bunch to fit them.

I don't know that Bronze Age Armenian will fit well with modelling the non-EHG "other" input into Yamnaya. It might do better than present day Armenian, but in the ADMIXTURE it is not like 100% teal or anything.

SimonW: This principle works also the other way round, it explains perfectly why South Asian IEs differ autosomally from their Andronovo ancestors.

Bear in mind that the samples we have are from the Northeastern far edge of Andronovo horizon, in Russia, I think - see the https://drive.google.com/file/d/0B9o3EYTdM8lQdzRlWXBQaXlnMGc/view?usp=sharing.

The distance from Sintashta to South Asia is quite high, and probably had many people in it, which is more obvious on an equal area map than the Mercator type projection that for some reason this paper has used - http://i.imgur.com/KdmfHsX.jpg.

Thanks Matt, I agree with your analysis. From the info in the paper it doesn't seem too likely that the BA Armenians have any European or even steppe origin, but it's also impossible to know if they will be a good fit or not for the non-EHG part of Yamnaya. It's just a possibility, but also the fact that the oldest one is from ca. 1800 BC doesn't help much. We will need older samples in any case.

First let's see how these ones look like when David can run them through a few tests to get a more accurate idea.

I don't think there was any ass whooping, but more likely that a good chunk of R1b went on to West Asia, leaving a hole for R1a to fill. I think that the story in Europe is R1b groups adapting to a more sedentary life with crops allowing their pop explosion to basically overtake what was more R1a heavy areas in NC Europe.

@ Alberto, yes, definitely older samples would be much more useful for this, maybe this is part of what Reich / Patterson's group is up to.

@ Krefter and others, I've taken the k16 (if I count correctly) onwards of the ADMIXTURE and rotated and labelled - http://i.imgur.com/yq5PjV2.jpg and http://i.imgur.com/hgKysyw.jpg, second of these has the Asian populations moved, as that might make it easier to compare between ancient and modern West Eurasians.

Btw, anyone have any thoughts about this Remedello Culture? It seems like the Baden Culture (CO1) sample reported in Gamba et al, where Gimbutas thought Baden should be Indo-Europeanised, apparently individual burials following an Indo-European pattern, etc. but at the genetic level it just looks more or less Early-Middle Neolithic. Same thing here, where Remedello was thought to be Indo-Europeanised on a similar basis, but look EN or MN.

Maybe hints at some disassociation with the "Kurgan" "package" (fortified hill settlements, weapons in burials, maybe even horses) which could spread culturally as compared to the steppe pastoralist type of way of life, which may have been more associated with Corded Ware and demic change? Even though CW was itself affected by this cultural change. Probably need more Chalcolithic culture samples.

Gimbutas thought IRC that kurgan type culture changes were connected with ideology and not movements of peoples, so perhaps that is a minor validation of her idea, even though at the same time, obviously we are seeing this quite large genetic change with CW and then BB and other Bronze Age Europeans.

I remember Anthony saying something about them possibly being more about funerary games, like chariot racing in Rome, rather than an actual weapon of war. I'm not sure if there was more war with other groups than within groups. The Bronze Age seems rather peaceful compared to the Neolithic, with all the headhunting still going on. The other option is still possible though.

* Chariots: Im sure a lot of it was about display and status. But surely it wasn;t all just posturing. One can fathom that the chiefs of Sintashta would have wanted to very much safeguard the gold mine (ie bronze mines) their forefathers had ventured.

* Matt & Kurganized Cultures. Your points are very true. That is why I always argued that CWC might have a 'native' European origin - and it still might in the sense that R1a does not appear to have arisen east of the Caspian steppe. I still cannot overlook the role of C-T culture colonizing the Ukrainian steppe (Neolithic culture or not), and hence the rise of European subclades of R1a there.

Whatever the case, my point is, Copper Age (pre-Yamnaya) cultures like CT, Baden, Cernavoda, etc looked "Kurganized' to Gimbutas but she was only partly correct. What they show is all the hallmarks of the secondary products revolution (c/- the Sherrats)- with a shift toward a more pastoralist economy, more dispersed settlements, individual burials, etc. So these features, as I had always maintained, had roots in the Late Neolithic cultures of Europe (as well as Western Asia) - even if west European R1b did not come from Baden/Hungary, but from the steppe. It is also pertinent that such cultures continued in Hungary, and Yamnaya appears to have had very minimal impact. North central Europe was far more demographically volatile, and hence R1b pastoralists made a far greater impact, as they appear to have been filling a vacuum, and not rampaging through with their (non-existent) equestrianism.

* it is a shame there is a lack of late of Late Neolithic samples from Caucasus (Nth and Sth) , and any from Ukraine. These are vital if we are to clarify 'the story of R1b'. There is always a remote chance that R1b - at least Z2103- spread from the Caucasus to Anatolia , Greece, directly, than via north (Ukraine & Balkans).

* I again highlight the floruit of J2 during the Late Bronze and Iron Ages. They appear in the proto-Scythian world (Sth Russia), the Carpathians (and presumably we will find them in Greece, Italy and Iran at this time) - areas we know the first IEs appear after the Middle Bronze Age "collapses".

I think proto-Corded Ware expanded from around the Don, probably the northern part, near the current Ukrainian/Russian border. The area I'm thinking of is marked by the black diamonds on the lower right map. Sintashta/Andonovo also came from there.

CT might have influenced far western Yamnaya and early Corded Ware people as they moved west. But we don't have any direct evidence of this yet. GAC may also have mixed with Corded Ware, but again, we don't have any direct proof yet.

Seems geographically plausible. But how do you suppose these Don groups developed & expanded their pastoral economy? Unless Khazhanov was completely wrong, they cannot have simply switched from foragers to pastoralists. Some kind of agricultural background is first required- it had to have from somewhere west or south. Cuacasian wives ?

For completing my discourse:Artur Martyka It's nice that you've quoted me and said that you've won (on Eurogenes). I haven't even known that I'm beaten.Gioiello Tognoni Did I understand well your post? It seemed to me that you recognized that Bell Beakers went from West and not from East, if CW were superimposed from them. Anyway I didn't say I won against you (whom I consider a friend and an open person), but against all who preferred banning me instead of confronting opinions. You are young. I am writing about these arguments from 2007. About one thing I disagree with you: all those old subclades of R1a you spoke about I think didn't come from CW, but are the witness that also R1a came from West... and we are waiting for the tests we spoke about a lot in the past.----------------------------------Of course I prefer your irony to the banishments of the past, and you are the owner of the blog. Honor to you. Anyway we are doing hypotheses at the light of the reason, but will be the facts to decide who is right and who is wrong. Don Quixote certainly is a relative of my R-Z2110 and the father haplogroups who migrated tio Iberia begining from 7500 BC as Zilhao demonstrated already many years ago.But what did happen to eng.molgen and YFull? The same people who like me?

For completing my discourse:Artur Martyka It's nice that you've quoted me and said that you've won (on Eurogenes). I haven't even known that I'm beaten.Gioiello Tognoni Did I understand well your post? It seemed to me that you recognized that Bell Beakers went from West and not from East, if CW were superimposed from them. Anyway I didn't say I won against you (whom I consider a friend and an open person), but against all who preferred banning me instead of confronting opinions. You are young. I am writing about these arguments from 2007. About one thing I disagree with you: all those old subclades of R1a you spoke about I think didn't come from CW, but are the witness that also R1a came from West... and we are waiting for the tests we spoke about a lot in the past.----------------------------------Of course I prefer your irony to the banishments of the past, and you are the owner of the blog. Honor to you. Anyway we are doing hypotheses at the light of the reason, but will be the facts to decide who is right and who is wrong. Don Quixote certainly is a relative of my R-Z2110 and the other haplogroups who migrated to Iberia beginning from 7500 BC as Zilhao demonstrated already many years ago.But what did happen to eng.molgen and YFull? The same people who like me?

@ DavidskiI wasn't able to delete my post with some mistakes. The second is the right one. About what you say, perhaps you know that I based all my theories about the Y and the mt, because more reliable. Autosome may say all what someone wants, and if my theory of the "Italian Refugium" against that of the kurganists will be demonstrated right, it will be demonstrated that I was right. I think that just about EHG and WHG have been made much confusion. All the tests about R1a-M420 and R1a-M459 are demonstrating that the origin was in Western Europe. Of course R1 may have come from Central Asia, but also about that I wouldn't be sure. Mal'ta boy is a dead line and also here it will be the aDNA to say who is right and who is wrong.

What I mean is that if the EHG groups around the Don and Dnieper acquired their pastoralism knowhow and southern admixture gradually, via long distance and small scale interactions, it might be very difficult to learn how this happened exactly, even with lots of ancient genomes.

On the other hand, if there were a few big migrations that led the process, like, say, major CT and/or Maikop incursions onto the steppe, then they'll be easy to uncover eventually.

I suspect that the Majkop chiefs are essentially native EHG guys. Partial admixture (viz the south) at best.If anything, they learned pastoralism and became 'acculturated" from the west (Balkans, Cucuteni, etc). I hope Niktin and his team are successful in their upcoming projects.

@ Mike ThomasI did know that, very likely I2a-M26 and the subclade close to Sardinians, where hg. I is Palaeolithic (if not directly in Sardinia, certainly in Italy). But I know many other things:1) Also in Yamnaya has been found I2a, and, if linked to the Italian one, it could be the witness of the supposed migration from Italy Eastwards also of the first subclades of R1b: I am R1b1a2-Z2110* with 94 mutations from Full Genome and have a link with a French Basque and Winterowd/Barnum. Very likely I am in Italy from 6000 years (as to YFull, 9000 as to MJost and me before him) and all the Eastern R-CTS9219 derive from my brother line CTS7556 born certainly in Western Europe.2) I am saying from years that not only Italy, but also my region (Tuscany), have different Y-s place by place: the 51 Tuscans of the 1KGP come from a little town near Florence: in Western Tuscany I found 100% R1b, from R-M269* to my R-L23 to R-U152, thus Remedello demonstrates only that they were descendants of the old Italian hunter-gatherers, but I am asking from years that samples from Rinaldone and the Arene Candide are tested. I think that Tyrrhenian Italy will be found plenty of R1b but also J, E, G, T from at least the Mesolithic and some hgs also from Palaeolitic.

@ Nirjhar007Of course we have in Italy many R1a subclades. Whereas I think that Italy was the refugium of R1b1-L389+ (I demonsttrated through Raza and Joshi that India has only old subclades come probably from Central Asia or nearby the Caucasus), I think that R1a* was in Northern Western Europe, even though Italy has samples of R1a-M420, of R1a-M459, and the unique sample of R1a-M420 found in Europe from Underhuill was found in Italy. But I think that R1a developed in Western Central Europe. It seems clear that Asian and Indian R1a came from Eastern Europe with the Indo-European satem languages. So Italy has many R1a subclades come later to Italy, not all recently: some 4000, 3000 or 2000 years ago, and they may be linked to each phase of the development of this haplogroup in Central Europe. The Slav invasion after the fall of the Roman Empire interested the Balkans, but not Italy, so I think that Italy lacks the most recent subclades of R1a linked to the Slav peoples.

I think not only the Armenian but also Mycenean Greek and maybe some non-Anatolian IEs in West Asia are related to Yamna R1b. Also a probable candidate is the Dacian a Paleobalkanic language. The routes of diffusion can be various trough West Asia for Greeks, or trough Balkans for some Paleobalkanic languages.It is possible that Greek was present on the South of Black Sea (later known as Pontus greeks) much earlier than we thought.

As far as I remember R1b is higher in Southern Greece and not in north. So a route trough Crete is probable. Later they move to South Italia.

ADMIXTURE in Allentoft is very simple, and they explain it well in the Supp. Info. The only reason to look at in detail is to see how ancient groups score compared to each other and modern groups, not see find what the pattern ADMIXTURE is showing.

Sintashta and Andronovo always score most similar to Corded Ware, but with slightly more Caucasus/South Asian and less EEF/Near Eastern. Bell Beaker, Unetice, Nordic LN/BA, and BA/IR Lithuanian/ score like North Europeans(Estonian, Russian, Norwegian in this case). Remedello Italy scores like EEF and Sardinians.

The last thing I want to investigate is LBA/IR Montengro. The only SE Euro reference in Allentoft's ADMIXTURE are Greeks. From quick glances it has seemed LN/IR Montengro scores similarly to Greeks but with slightly less EEF/Near Eastern. IR and LBA Montengro seem to score differently, with LBA scoring higher in EEF/Near Eastern.

I forget to mention Albanians. Especially Kosovo Albanians for whom the R1b is the second after the E1b1. So them also can be related to that Yamna spreading R1b.What are the subclades of Albanian R1b?? Do they match?

Gioiello,Thanks For your Suggestions:-), On Indo-Iranian i wait to see aDNA from SC Asia etc before making any kind judgement which is the wise thing to do.David, Thank you for your short suggestion also:D.Aram, A thought The Armenian Iron age J2b recalls the frequency of J2b among Greeks, who, as known, have a language close to Armenian. Maybe that was really the arrival of a Phrygian people who brought the Armenian language? what do you think?:).

I think Kurti tested that J2b. It seems he looks more Iranian like and not Balkanic like. So this J2b could be related to Indian J2b :) In modern Armenians we have two types of J2b. The half is Balkanic type the half is the Eastern type (that is present in India also). But overall this J2b is minority in Armenia. The majority is J2a.

Now the most perplexing thing is the origin of that Yamna R1b. If it is a result of founder effect then it could come from everywhere. Also the late appearance of J2 is an interesting phenomena. This mean that the debate will continue. Between the Steppe and a more Southern origin of IE. ;) I think the Z93 will be decisive.

I forget another issue. The "teal" component. Even if Z93 comes from North the issue of teal component will remain, because it seems omnipresent in all BE related cultures. This "teal" cannot come Cucteni-Tripolye or Europe. So maybe Maykop or South Central Asia. And how this teal relates to the IE problem.?

@Nirjhar007Of course you are right in waiting for aDNA tests in Asia. This is always the decisive proof, but was I to say that an old R1a in India seemed very old for its STRs values and asked the analysis of Klyosov, which didn't come, and many of us supported two migrations of R1a to India: one old before the Indo-European one of the Second Millennium BC. Thus everything is possible.But to @Aram Palyan ("What are the subclades of Albanian R1b?? Do they match?") I have to say that:1) Albanian R1b is carachteristic for an R-L23 subclade (the name "Balkan cluster" is mine and I was the discover of it with Argiedude): now we know that this subclade is very recent, linked to Iberian haplotypes, descends from CTS7556 come very likely from Western Europe and brother clade to my R-Z2110 in Western Europe from 6000 or 9000 years.2) There are also R-M269, and so far the most part of subclades come from Western Europe with only a tiny subclade (Mehmet Laz) in Turkey, and the "Jewish cluster" has perhaps its origin in the Greek world if the ancestor of Anne Hart accpted to do a deep SNP test.3) About the origin of R1b1-L389* I have written tons of letters from many years.

@ Mike T: Whatever the case, my point is, Copper Age (pre-Yamnaya) cultures like CT, Baden, Cernavoda, etc looked "Kurganized' to Gimbutas but she was only partly correct. What they show is all the hallmarks of the secondary products revolution (c/- the Sherrats)- with a shift toward a more pastoralist economy, more dispersed settlements, individual burials, etc. So these features, as I had always maintained, had roots in the Late Neolithic cultures of Europe (as well as Western Asia) - even if west European R1b did not come from Baden/Hungary, but from the steppe. It is also pertinent that such cultures continued in Hungary, and Yamnaya appears to have had very minimal impact.

I don't have the knowledge of archaeology to sense check this, but if I have understood correctly, it does sound possible to be thinking of these features as arising in parallel to a similar material stimulus, among Neolithic European and steppe societies, even if there was still also a large steppe movement to North-Central Europe, who would have brought much of their culture with them.

Re: continuity of local cultures in Hungary, as you have knowledge on this, do you think this paper tells us anything about the transition of cultures there? For Hungary, they have samples from Vatya, Maros, "MBA", and Vatya culture contexts all around 2200BC to 1600BC.

The Haak paper suggested the Bronze Age Hungarians had some autosomal (demic) impact from Yamnaya, but not very much, similar levels to the lowest of the South East Europeans today. D statistics here have not quantified the input but also agree that the BA Hungarian input from Yamnaya was relatively low compared to Bell Beaker (who are lower relative to Corded Ware).

Looking at the D statistics in the supplement, of the form Y(Yoruba,Yamnaya;Bronze Age Hungarian,other ancient), there is a larger difference between the Bronze Age Hungarians and Bell Beaker than between Bell Beaker and Corded Ware, even though BB and CW were estimated by Haak to differ by around 25% Yamnaya input (from 75% to 50%). So it seems possible that once these samples are put through the qpAdm methods from Haak, they'll have a similar 10%-20% Yamnaya only as with the Hungarian Bronze Age samples from Gamba tested by Haak.

Does this fit with what you'd have expected?

Re: CT and Yamnaya, if I understand correctly, it seems like you could have a cultural transfer from CT->Western Yamnaya->Eastern Yamnaya (who we have samples for) with ultimately little genetic transfer between CT->Eastern Yamnaya at the ends of the scale (because there doesn't seem like there would be much). I guess that is more of an archaeological question.

The 25% north euro in copper age Armenians might mean R1b metal workers from Yamnaya with 50% north euro moving into some part of the Armenian highlands that had copper and marrying into the locals hence becoming 25% north euro in the region where they mixed. The reduction over time could then be further dilution with the wider original population.

"they cannot have simply switched from foragers to pastoralists. Some kind of agricultural background is first required- it had to have from somewhere west or south. Cuacasian wives ?"

One straightforward way it could have happened is if foot herders encroached on the territory of mounted HGs and the HGs won the ensuing battle. Something like this could potentially create a warrior aristocracy type culture automatically.

Alternatively trading horses or copper for wives and the females brought pastoralism with them.

HGs transforming to a warrior elite over a herder base without themselves ever going through a herder/farmer transition makes a lot of sense imo.

although after some quick reading some of the history another possible explanation for a decline in the north euro element may be all the wars that came along after and getting pushed away from the Lake Van area.

Compare this to modern Armenians:North_Sea 4.05%Atlantic 2.34%Baltic 1.06%Eastern_Euro 2.01%

Obviously all of these rather north European components are higher in the Bronze Age Armenian.That's why the oracle chooses mixtures of Georgians + various North Europeans as the best approximations. Georgians are one of the purest, most typical populations of the Caucasus, as West_Asian and Caucasus components of various calculators repeatedly peaked there. So the finding that Bronze Age Armenians are like Georgians + North Germans, Scots or Danes means something. It's crazy to suggest that Bronze Age Armenians are a regular, pure West Asian population without substantial Yamnaya-related admixture.

Of course, this is just the „old Eurogenes K15“, not the more recent K8 and K9, but the existence of the latter do not at all invalidate the K15, which is still a great tool, IMO.

But you can also check Dodecad K12b by Dienekes:Bronze Age Armenian RISE423 has 22.57% of the North_European component. The various modern Armenian samples have 3 – 4.5% of this component. Of course these figures are slightly distorted by the calculator effect, but it's still more than obvious that Bronze Age Armenians were substantially admixed with a Yamnaya-related population.

"It's crazy to suggest that Bronze Age Armenians are a regular, pure West Asian population without substantial Yamnaya-related admixture."

I'm not suggesting they're regular West Asian population. I'm saying that if they turn out to be some 13-15% WHG in K8, it will match the "Caucasus-like" population that entered the steppe, because that population had such amount of WHG. That Caucasus-like population was absolutely not a pure West Asian population. But not because of the 13-15% WHG (which could be perfectly normal for Anatolia since the Neolithic), but because of some 32% ANE (EEF had 0% of it).

I have no idea if these Bronze Age Armenian samples will anything close to that ANE. I have to wait and see.

But I'm telling you that it's early to jump to conclusions, one way or another. Matt explained in detail the stats from the paper, and you can look at them on your own. They do suggest a rather different thing that the calculators based on modern populations, which might be dealing with an odd population for them and having troubles giving accurate results.

I'm being cautious for a reason. And I think we should all be. I'm not saying what you are suggesting is necessarily wrong, but it's too early to give it for granted. Let's wait and see.

For example, in K12b we have Samara_HG at 74% North_European. And we have Yamnaya at 60% North_European. This means that the population who mixed with EHG in the Samara region should have some 45% North_European, right?

So again, let's wait a few days to see what those BA Armenians really are.

I think the Armenian E1b sample is potentially as significant as the R1b.

If it's representative of the base population which an intrusive R1b population mixed with (and it might not be) and if that E1b represents a population layer that was mostly displaced outside of a few mountain refuges (and it might not be) then a population made up of E1b people + R1b people is going to be quite unusual.

I'm sure there must be some places with a lot of E1b and R1b but I can't think of any offhand.

@ AlbertoWell in that sense, Georgians are not a pure West Asian population either, because they've got ANE and teal. They are pure and typical for modern West Asians, i.e. score high in modern West Asian and Caucasus components. And yes, it's still quite likely that some ANE flowed west along the South Caspian. Because, as you said, the Caucasus/West Asian people who admixed with EHG on the steppe had some ANE. But in contrast to you I have quite some confidence in this oracle stuff, at least if used with sensible admixture calculators. If Bronze Age Armenians were without any steppe-related admixture, I would expect them to be come out similar to Sardinians + Tajiks or Kalash or something like that. That West + ancient south central Asian ancestry would by chance result in affinity to North Europeans seems rather miraculous to me. But we don't have to come to an agreement, you're free to wait for more analyses.

Gioiello,Yes I also remember similar suggestion from a certain Stepanov et al. but i'm not aware it was published or not But i do think rather than modern STRs we should see the Specific SNPs of Ancient DNA and then Make conclusions.It is also quite interesting that none of the Yamnaya samples have produced any R1a but CWC have and also the late bronze age samples of Central Asia i'm actually trying to find more about the Connection between Central Asia and CWC, i am suspecting the R1a origins maybe either in C Asia or NC Europe!Of Course Everything is possible but not if you follow the Kurgan Hypothesis;)Aram, The Teal Component is certainly a curve ball if one wants to postulate NC Europe as PIE land its sudden presence in Europe can only be Linked with IE Progress i think...

@ SimonW, yes what Alberto says about how the BA Armenians may be an odd population for the confines of the components that Eurogenes K15 based on the modern variation. Which might not be that fit to represent them.

They might have had a population input from Yamnaya, but I don't think they will be anything like 85% Chechen plus 15% North European or anything.

To be repetitive, look at the f3 statistics for f3(Ju_Hoan_North,Test,Yamnaya), lower to higher:

If f3 stats are anything close to linear (and I think they should be), then using that figure for RISE407 of 85% Chechen plus 15% Swedish and Norwegian as a proxy for Swedish would be (0.22952703+0.0412677) = 0.270794. A Bronze Age Armenian would be noticeably closer to Yamnaya, than Georgian. Instead it is very similar to Georgian in how close it is to Yamnaya.

BTW keeping aside the vision of future past of yours;), i certainly think that Indo-Iranian BMAC metallurgists went there in Sintashta area, so it will be interesting to see what Y-DNA Mutation BMAC aDNA samples provide....

Note that non of the Bronze age Scandinavian Y DNA is R1a, and 2/3 of Late Neolithic ones are(Including a Battle Axe Swede from ~2800BC). In Allentoft's ADMIXTURE Late Neolithic Scandinavians score higher in Caucasus/South Asian than Bronze age ones. The Late Neolithic ones look somewhat Corded Ware-like and the Bronze age ones look modern.

It's strange how downstream some of the ancient Y DNA is. I didn't expect Sintashta to find a downstream subclade of Z93, or Bell Beaker a downstream subclade of P312.

"BTW keeping aside the vision of future past of yours;), i certainly think that Indo-Iranian BMAC metallurgists went there in Sintashta area, so it will be interesting to see what Y-DNA Mutation BMAC aDNA samples provide...."

One analysis of Sintashta and Andronovo showed high ASE and low WHG, and another looked NorthEast European. Some at Anthrogencia suggested BMAC had influenced Sintashta, which can explain their ASE. I think Sintashta was most like Corded Ware(based on Allentoft's ADMIXTURE), and that's where their R1a-connection comes from. But I'm just waiting for better anyalysis of Sintashta/Andronovo.

I agree, I think Northwestern European R1b is mostly from a Corded Ware-derived founder effect.

Here is what is known.

1. Out of 12 yamnaya samples 11 have come back r1b and none r1a. These samples come from the extreme northeast of the yamnaya horizon and the south central region. While geography will likely relate to the diversity of yamnaya ydna, we should not forget that yamnaya itself, i.e. the culture, is a predictor of ydna. Thus far, the correlation between culture and genetics has bore out through the consistency of yamnaya sample results. Also, we should not arbitrarily divide the yamnaya horizon based simply on what has and hasn't been sampled then assume the unsampled regions are somehow expected to be different.

2. The yamnaya inhabited a geographic region, the steppe, and they were a very mobile people. It would be extremely strange for a significant portion of western yamnaya to be dominated by r1a while all of those sampled outside the west have completely lacked r1a. Outside of the yamnaya culture, the PC steppe have also produced 1 mesolithic sample of r1b and 1 r1b sample contemporary to yamnaya while not being labeled as yamnaya. It appears that around the time of yamnaya and even before, there was a strong presence of r1b on the PC steppe.

3. northwest europeans cannot be derived from corded ware on account that corded ware hardly had any direct presence that far west and because central europeans, the only possible link to CW have less yamnaya heritage. The large amount of r1b found at an early stage of the eastern bell beakers from germany, Cz, and Hungary does not bode with a recent founder effect, but its the autosomal evidence that disproves a founder effect. The migration has a large base, not a small corner.

4. The yamnaya themselves expanded into hungary. You seem to have completely ignored this well known fact. These hungarian yamnaya derive from western steppe yamnaya, and its almost certainly a source of r1b for western europe, rather than the r1a heavy CW.

If your interpretations need motivation, remember, its northwest europeans who's r1b MUST derive from the yamnaya. R1a in corded ware need NOT have come from the yamnaya..

Based on f3 for Sintashta, WHG, Malta and Neolithic Central Europe - http://i.imgur.com/ZedsV1w.png

I circled the Bronze Age Armenian, and you can see that on these PCA it's generally pretty close to the other West Asians (Georgians, Abkhazians, Turks).

Considering only WHG and Malta outgroup f3, BA Armenian is in the same place as West Asians.

However there is more separation from them when the Neolithic Central European f3 comes into play, and they are pushed away from other West Asians.

Iron Age Armenian in contrast seems pushed more towards the Balkans and Southern Europe relative to Bronze Age Armenian and present day West Asians (some genetic influence via the European Balkans), and finally modern Armenians are somewhat "south" of them both (some genetic influence from the south). That all seems plausible given the history of Armenia? I don't know though.

A weird thing on these is that the Unetice and Bell Beaker seem to have less of a WHG / north direction than I would have expected from Haak, perhaps this is because of their differences in sampling location?

Isn't it surprising to find that R1a-Z2124 in Sintashta? If these people were a recent arrival from Europe (as they seem to be), shouldn't that mean that the 3 mutations upstream from it (Z94, Z93 and Z645) originated in Europe and therefor Europe should have a lot of it? Or am I misunderstanding the sequence?

Thanks for those PCAs. They do seem to give a much more clear vision of those BA Armenian samples. It seems to me that they could be related to the population that entered the steppe, but after 1500 years in the area they became quite more West Asian. The "original" population should plot more around where Tajiks are, I'd guess.

On a different note: Who is that Stalingrad Quarry sample? He is contemporary with Yamnaya and not from far away from where the Yamnaya samples were taken. Why is he not labelled as Yamnaya? There must be a good reason, because he does plot in a different place. As if instead of being 50/50 EHG/Armenian-like he was about 75% Armenian-like. Anyone knows to which culture he is related to?

I see. Well, I think that the European ancestry should be proved. Matt's PCAs above also plot Sintashta near CW. So maybe it's a mistake in the identification of the haplogroup, yes. We'll have to wait for confirmation.

Geneticker is updating is Y DNA analysis with files that have calls in more Y SNPs. All the P1s, K2s, etc. from Corded Ware are becoming R1a-M417, although I guess they didn't get a call in Z283 or S224.

BTW, Late Neolithic Scandinavian Y DNA so far is 2/3 R1a-M417 and 1/3 I(I1?), and a low coverage sample. Bronze age Scandinavian Y DNA is 2/3 R1b-M269 and 1/3 I1, and a low coverage sample. In Allentoft's ADMIXTURE Late Neolithic Scandinavians score similar to Corded Ware and Bronze age ones to Bell Beaker.

"With some further investigating and team work it has been determined that the haplogroup J2a sample from Iron Age Altai belongs to the LM410 subclade. Some further digging and reading also revealed that this sample is labelled RISE602 in the RISE project and was discovered in the Sary-Bel kurgan. It has been dated to Iron Age (900/700 BC-AD 500/1000 ) and a rough translation from a Russian site has this to say about this burial site. ''So Sarah burial cemetery date back to the beginning of Bel-Hun-Sarmatian time and,may belong to the people, which penetrated into the Altai Mountains to the southeast"."

"The Bulan-Kobin culture evolved out of the Pazyryk culture which was Scythian. So there seems like there is a good chance then that this J2a-L24 male may have belonged to the Bulan-Kobin culture which had its origins in the earlier Pazyryk culture.

Apparently the other J2 sample from Iron Age Russia actually also comes from Altai. It comes from a site called Kytmanovo. So this is now two J2 samples from Iron Age Altai. This is getting very interesting. Perhaps it was J2/J2a men that introduced iron technology to this part of the world. "

I said it sooo many times now it is finally proven. J2a beside R1 and R2 and some G is one of the main Haplogroups of the ancient Indo_Iranians.

A person just needs to look at the overall distribution of Haplogroups among modern Indo_Iranian speakers. J2a is among the most frequent Haplogroups in ALL Indo_Iranian speakers.

Rather native to this land. What if (more likely) a Southernwestern input(Semites) changed the genetic landscape.

It can't be a "Northern input" because this Bronze Age Armenian sample has typical Farmer DNA for most. And their Northern Admixture is on pair with that of North Caucasians. Now let us take in mind that the people living in Armenia back than were Paleo_Caucasians and Indo_Iranians.

"I meant north European-like admixed, not in a geographical sense northern. I won't put any money on a concrete origin. But the exciting thing is that, judging from this evidence, R1b in Armenians really came with the Armenian language and from steppe-related people. That is, the R1b-L23 there didn't originate in the South Caspian or in West Asia, and the same holds true for the Armenian language. Conversely R1b-L23 in Yamnaya was from EHG, not from the West Asian element."

The Armenian language reached Central Anatolia by 1200 BC at best. Those results predate the arrival of the Armenian language by over 1000 years. I don't see how R1b could have come with the Armenian language. What about V88 in the Levant and Neolithic Spain? The results of all the ancient Armenian samples look like a mix of North Caucasian, West European and Iranic.

Armenian R1b is bottlenecked but Iranic/Kurdish R1b is diverse with allot of basal m343.

"It's fascinating to see how the North European-like admixture initially, in the Bronze Age, was quite substantial (about 25%), then got weaker in the Iron Age (17%) and is very weak nowadays. "

Yes because of the Semite expansion (Assyrians) into the North we see how Red Sea/EastMed type ancestry rises and how North Caucasian/Iranic type ancestry decreases. It has nothing to do with an "Indo European" expansion, because again the oldest with the most North European type ancestry predates any known Indo European language. This "NOrth European" like ancestry mus be native component of this region.

"Apparently they mixed more and more with the locals but were able to preserve their language and ethnicity nonetheless."

Absolutely not, how can someone come to this conclusion if all samples are this way. If all samples are different that means THEY ARE THE LOCALS and got admixture from South which is historically evident. Their North European type ancestry looks pretty much in the range of what modern North Caucasians have. Hurro_Urartians (Natives) are believed to have spoken a Northeast Caucasian/Indo_Iranic related languages.

"It has been observed via physical anthropology that a long headed "Nordic" looking type entered Armenia in the late Bronze Age. Modern Armenian crania differ a lot from this. So maybe this is one of the few times where considerable changes in cranial metrics were paralleled by genetic changes?"

physically Nordic from a predominantly dark haired and eyed population such as Yamna at a time when there is no sign of any Yamna movement towards the region (2500 BC)? I don't think so

exactly, "European" admixture such as Atlantic_Med and Caucasus lacked in Yamna samples, but are very high in Bronze Age Armenian samples during a period when there is no known migration from North to anywhere in Western Asia. The 20% North European are in line with the North European ancestry in North Caucasian.

If anything those guys are the natives as seen on all the samples and a southern expansion started and changed the genetic landscape.

I don't understand how some people have so hard time to accept that some "North European" type ancestrs (which is WHG and ANE whatsoever) can be native to the region, but at the same time have no hard time in accepting that those "Gedrosia_Caucasus" genes in the Steppe are probably native to the region anyways. So why does it work one but not the other way?

Maykop is a culture which spans from Northeast Anatolia to Northwest Caucasus. From the wording of the paper it seems that Maykop had substantial EHG ancestry. Why is it so hard to accept that EHG type ancestry could have been in the region since long time?

If so, what do you think the chances are that some Y-DNA J2, or perhaps J2a lineages specifically, left the Near East during the collapse period and that is why we see it appearing in Hungary around the collapse period and on the steppes during the Iron Age?

I remember that the J2a lineage from Bronze Age Hungary was found in a Urnfield context and was dated to around 1100-1270 BC (around the collapse period). I also remember reading that the cremation rituals of the Urnfield culture were rooted in traditions from the East Mediterranean.

Geneticker has made updates. The I1 from LN and BA Sweden is I1a3a1. That's pretty downstream. I wonder if most I1 is from Funnel Beaker. The J from LBA Armenia is J2b2f, and apparently there's another E1b1b1 from Armenia. Also, LBA Lithuania is R1a1a1.

I've always thought it odd how there were so many references in classical writing to red haired populations over such a wide area: China's western barbarians, Scotland, Libya, Thrace, Germany etc and it seemed to me it was too much of a coincidence to not be a flashing neon clue to something or other - just a question of finding out what it was a clue for.

I used to think those written references might have meant that red hair originally had a very widespread distribution which had then shrunk over time. I now wonder about another possibility i.e. the source region for red hair somehow got tied up with something else that spread dramatically over a very wide area and the red hair went along for the ride.

If copper working first started in the southern Urals (or even if it started elsewhere but copper workers moved to the Kargaly region quite soon after for mining reasons and recruited locals) then that might have created an association between red hair and mining and then when a new source of copper was found where would you look for miners?

It would be an amazing fluke but if correct would create a kind of barium meal for the spread of neolithic mining. In most places it would be a hidden clue because it's recessive and the percentage of miners in the total population in most places would be. It would only show up visibly in regions where the miner dna was a significant percentage for example regions where the pre-existing population density was very low.

It throws an interesting light not only on questions like why are Scotland/Ireland and the Udmurts the reddest with a huge gap in between but also things like King Tut being rumored to be ydna R1b and red haired.

Copper workers in ancient Egypt are likely to have been wealthy and could conceivably have married into the local elite.

If the R1b/copper thing does turn out to be a thing then it seems likely something similar may have happened with Bronze/Iron also i.e. people from particularly favored source regions spreading along the trade networks.

They're West Manych Catacomb. That explains the I2 in Hungary. It's Catacomb derived. There should be R1b there too. Whether it's Z2103-7, or L51 is another question. I have a feeling L51 will end up being found in Corded Ware, at this rate.

"The large amount of r1b found at an early stage of the eastern bell beakers from germany, Cz, and Hungary does not bode with a recent founder effect, but its the autosomal evidence that disproves a founder effect. The migration has a large base, not a small corner."

It's not a large amount though; it's a large percentage of a small amount and if it represented a metal working caste among a pastoralist majority maybe they were buried differently?

(sky burial for the patoralists and back into the earth for the miners?)

I agree the distribution of R1b and R1a in Europe (mixed together but the proportions almost mirrored west/east) doesn't seem to make make sense unless the options are

1) R1b from the west, R1a from the east2) R1b (pushed?) west en masse by R1a3) A boomerang where R1b went west in small numbers, expanded in situ and then moved east again in large numbers

and as there's no old R1b found in west (at least yet) we can scrub option 1 for now.

So it seems to me the logical possibilities given the odd distribution is either a mass R1b migration to the west or a boomerang where a small R1b migration went west and then exploded dramatically on the northern and western periphery of Europe for some reason.

(I appreciate that on the face of it your option seems the most plausible.)

* "I don't have the knowledge of archaeology to sense check this, but if I have understood correctly, it does sound possible to be thinking of these features as arising in parallel to a similar material stimulus, among Neolithic European and steppe societies, even if there was still also a large steppe movement to North-Central Europe, who would have brought much of their culture with them."

Yes; pastoralism can only have arisen where agriculture had already existed- ie Balkans, Anatolia, perhaps the fringes of the steppe near Caucasus and abutting the Balkans/ Romania. The Yamnaya movement into europe thius can be thus thought of as a continuation of a dynamic settlement cycle, whic brought their specific ideology pattern- wholly so in eastern hungary and nth Bulgaria; partially in corded ware.

* "Re: continuity of local cultures in Hungary, as you have knowledge on this, do you think this paper tells us anything about the transition of cultures there? For Hungary, they have samples from Vatya, Maros, "MBA", and Vatya culture contexts all around 2200BC to 1600BC.

The Haak paper suggested the Bronze Age Hungarians had some autosomal (demic) impact from Yamnaya, but not very much, similar levels to the lowest of the South East Europeans today. D statistics here have not quantified the input but also agree that the BA Hungarian input from Yamnaya was relatively low compared to Bell Beaker (who are lower relative to Corded Ware)."

The EBA Carpathian basin was a mosaic of different cultures. We need more samples; but we have all noted the present but low Uamnaya impact genetically.

*""Looking at the D statistics in the supplement, of the form Y(Yoruba,Yamnaya;Bronze Age Hungarian,other ancient), there is a larger difference between the Bronze Age Hungarians and Bell Beaker than between Bell Beaker and Corded Ware, even though BB and CW were estimated by Haak to differ by around 25% Yamnaya input (from 75% to 50%). So it seems possible that once these samples are put through the qpAdm methods from Haak, they'll have a similar 10%-20% Yamnaya only as with the Hungarian Bronze Age samples from Gamba tested by Haak.

Does this fit with what you'd have expected?""

Yes it does. Keep those stats coming. There is significant continuity in west Central Europe. And I'm still not happy with the Yamnaya & CWC analysis at present. We need pre-Bronze age samples from Poland, Baltic etc to level out what I suspect is an overestimation of Yamnaya migration to Northern Eastern Europe and the Baltic, perhaps by 20%?

*""Re: CT and Yamnaya, if I understand correctly, it seems like you could have a cultural transfer from CT->Western Yamnaya->Eastern Yamnaya (who we have samples for) with ultimately little genetic transfer between CT->Eastern Yamnaya at the ends of the scale (because there doesn't seem like there would be much). I guess that is more of an archaeological question.

I think CT would have impacted incipient west Yamnaya . West and east Yamnaya would have then exchanged and "levelled out", etc

"pastoralism can only have arisen where agriculture had already existed"

Disagree with this and think it may be the key to the whole thing.

Seems to me the minimum you'd need would be sedentary HGs with a lot of food and a confident, curious species capable of self-domestication e.g. goats.

Say for example you have a valley full of fruit trees and the local HGs become sedentary as a result.

All you might need for self-domestication is for those static HGs to have enough food to produce a lot of scraps and an animal confident enough to hang around the human sites feeding on the scraps.

Even easier if both the goats and humans got drunk a lot on fermented fruit from the fruit trees.

The thing about this idea is if a population developed goat herding before crop raising then for the time they were like that they could have been incredibly mobile and spread all over the place very rapidly - only figuring out agriculture later at the sites they spread to which were most suited to that e.g. the ones where there was naturally a lot of wild wheat.

I think someone dealing with such a complex issues needs always check himself for consistency. So before mixing linguistics here just a plain question. In Your model from where did Armenian R1b come from and in what period?

What R1a clades do the Chuvash(north of sintashta) exhibit. My impression is that they have european clades z280 along with finno-ugric markers for both y and mtdna. Its likely the area where finno-ugrics picked up indic loan words.

kazakhs on the other hand like the sintashta sample probably have z93 and r2 like south asians?

"pastoralism can only have arisen where agriculture had already existed"

Disagree with this and think it may be the key to the whole thing."

Grey. It doesn;t matter what "you disagree" with. I, as othersm have told you to stop pulling your little 'pet theories' out of your backside, and read some literature, instead of coming out with your banal cliche after cliche ("domino effect", "boomerang effect', and fetish for migrating copper simths).

"I think this lends support to the notion that Y-DNA R2-M479 represents a more southern phenomenon than R1-M173, which (per aDNA) appears ubiquitous across the steppe. I suspect we'll find some R2-M479 (including extinct lines) in aDNA from territories immediately below and including Kazakhstan and Kyrgyzstan from the paleolithic onwards."

There's absolutely no sensible reason to assume R2 has anything to do with any of these steppe movements. Some branches of R2a-M124 (L295-) may have played a secondary "piggyback" role in the spread of Indo-Iranian, but R2a-M124 in itself is too old, too "deep" in its' phylogenetic clustering and too southern in distribution.

I am fascinated about all these highly interesting findings. The area of proto-IE language(s) looks like being pretty northern. It is interesting that there is still no R in Hungary and neither in Italy in this paper. It is also fascinating if the Etruscans will turn out to be like Remedello guys and spoke possibly a language related to the language of Neolithic Hungarians as suggested by somebody on Eupedia forum. Anyway, it looks like IE languages came to Italy with R1b and later than expected (with Urnfield culture?). If Indian R1a is really Sintasha derived, that is also a kind of a bomb to many of us.

As for Uralic groups, this paper gives mostly only indirect or negative evidence. The area where most of the Uralic languages are spoken was unsampled. However, I take note that there is again no N in South Siberia. On the basis of the structure and age estimations of N1c and N1b, it looks like they arose in the area east of Belarussia (N1c) and in the Volga forest area (N1b). The area where N1c may have arisen is very near the area where David suggested the proto IE language developed.

@KristinaWe all expected Indo-Iranian R1a in Sintashta or Andronovo ,there is nothing surprising about that and it is the final end for all kind of OIT fantasies.

Finno-Ugrians are poorly understood people and further research is bitterly needed but both Finno-Ugrians and PIEs are more or less derived from Siberia if we go enough back in time and the structural similarities between PIE and Finno-Ugrian languages are maybe showing very ancient ties.

Aram Palyan, Although the scenario you propose on the whole appears to make sense, I have to disagree about R1b moving from Anatolia westwards through Crete. Archeologically there is evidence for movements of north Anatolian people to Greece at the turn from the early to the middle Bronze Age, but this influence didn't reach Crete. And afterwards the Minoan civilisation with its non-IE language flourished – so if the Anatolian intruders were IE, it's in line with our expectations that they didn't reach Crete. I think it's true that R1b now is more common in southern Greece than in the North, and likewise J2 is particularly common on Crete. But this pattern may have been shaped by the Medieval Slavic movement from the North, and anyway we shouldn't equate modern Cretans with their Minoan ancestors. Modern Cretans are ethnically Greek, and they also differ physically from the Minoans, among other things they are much taller.

@ SimonW, they're the paper's f3 stats from "Supplementary Table 11 (748 KB) - This table contains “Outgroup” f3-statistics for all combinations of ancient and modern groups; Human Origins dataset" - http://www.nature.com/nature/journal/v522/n7555/extref/nature14507-s6.xlsx. I don't have any means to run them myself.

I expect that they are the averages of groups and like you probably suspect there may be some heterogenity in the group.

Using their significance test Bronze Age Armenian comes out unadmixed for these, as does Yamnaya, while the Bronze Age Europeans and Bronze Age Steppe generally show the expected patterns are. Although Sintashta doesn't show admixture while Corded Ware and Andronovo do.

For Corded Ware, Bronze Age Armenian oddly provides a lower (stronger) admix signal with WHG and SHG than is found for Neolithic Farmers plus Yamnaya, which may be do to with either the lack of Middle Neolithic samples in this set (no Neolithic North and they're all early Neolithic) or a preference for simpler mixture combinations / tree structures of particular kinds in these f3 stats, which can be affected by post admixture drift.

Also true of Corded Ware. WHG plus Bronze Age Armenian is the strongest f3 for BA Hungarians and Corded Ware.

Strongest for 1000G Indians with a decent SNP overlap is of course Bronze Age Armenian with HanB (lacking any other references).

Armenian is certanly not related to the Anatolian branch. So how can a 2500 BC samples from Armenia have had any known Indo European admixture? That is blatan speculation with no value. Ancient historians point to a relation of Armenian and Phrygian as languages. Most linguists see a connection between Phrygian and Armenian. I doN't see how Armenian language could have been the R1b bringers to the region. Especiall if Bronze-Iron Age Armenia was populated by Hurrian_Urartaen and Indo Iranian type people and we know Hurro_Urartaen at least partially a language related to modern Northeast Caucasian to which also Lezgian belongs. And guess who have the highest frequency of R1b in the North Caucasus.

Kristiina,// If Indian R1a is really Sintasha derived, that is also a kind of a bomb to many of us.//What do you know about bomb:D? Just for some samples from Indian aDNA.DMXX,I think you are R2? anyway I Think its a bit like I which didn't move much....

I agree with you. Linking Bronze Age R1b specifically with Armenians seems hypothetical, but is of course possible. Linguistically I guess, Id expect Armenian to be much younger. But one cannot overlook its links with Iranian. So if Iranian and Indic were already present c. 1500 BC in the region, so to could have pre-proto-Armenian.

That is the reason why I am specifically speaking about the Armenian LANGUAGE.

Note language and genetic ancestry is not 100% the same.

Modern Armenians are a mix of pre Armenian locals and Proto Armenian speakers.

Maybe a 50/50 mix.

And the Indo Iranian substrata can be explained with that.

I am not trying to "Disprove" an native origin of Armenians. 1200BC in Anatolia is still BY FAR older than other languges in the region (Semitic, Turkic )

I am just trying to make sense and see that R1b couldn't have come via the Armenian language speakers. Also because R1b among them is bottle necked. Diversity of R1b is by far larger among the Kurds and some other Iranic speakers in the same region. But also among Lezgic speakers.

Skilur,I don't know if you know what logic is or not but certainly when you say ''We all expected Indo-Iranian R1a in Sintashta or Andronovo '' its Asian R1a we are talking about and IMO it don't have European Origins unless aDNA from there says otherwise, anyway we still don't know the exact SNPs for sure yet...

@ KristiinaThat Indian and Asian R1a (with the IE satem languages) were derived from Eastern Europe is perhaps a surprise for most people who preferred banning me instead of reading carefully my writings: I am saying that from many years (and continue to believe that R1a descends from WHGs).About the presence of R1b in Italy if you were right it would be the disproving of all my theory of the Italian Refugium, but I'd wait tests in Western European Bell Beakers and in Tyrrhenian Italy where I live with my R-L23-Z2110* linked certainly with Western European people and not with the samples found in Yamnaya. That Etruscans descends from Remedello and peoples linked to Rhaetians and Camuns is possible (and I am saying that too from the beginning, in spite of many blatanting Anthrogenicians). I don't believe in my compatriot theory (Mario Alinei) about the link of Etruscan with Hungarian, came lately from Central Asia and far from other Uralic languages certainly older in Eastern Europe, if not in an older link at the Nostratic level. Alfredo Trombetti said that Etruscan was intermediate between Caucasian and Indo-European and very likely the I2a-M26 in Italy spoke a Caucasian language which were possibly the Sardinian ones.

I don't speak language. I asked You about R1b in Armenia. Where did it come from? They are subclades of Z2103. I never said Armenian R1b is older or it is a source of someone else. I think You know the map of Z2103 in West Asia and Balkans. Who are this people in Your opinion that are present from Balkans to South Caspian?

Gioiello ,Eastern Europe was not the origin point of R1a-Z93 because its too diverse in Asia its like saying Z-283 is West Asian in Origin since some population shows it.... We need aDNA from southern regions and also earliers time from SC Asia and C Asia to testify the origins of R1a...

@ Mike ThomasUnfortunately I am writing with a little laptop, very slow... Why I think that R1b1-L389* was in the "Italian Refugium"? I wrote about 10000 letters about that, from 2007 on Rootsweb till Anthrogenica which dedicated me a thread "Rathna's assessment of genetic material" before banning me. That threas has been visited from pretty 50000 times. It is in the General, lane 9, difficult to find but many people contine to visit it. It is useless now that I repeat all my arguments: now we are at the decisive proof: the aDNA. If it will be found I was right, if not I was wrong. The same is worth for Nirjhar007, who I already said he has all the right to wait for aDNAtest in India or Souther Asia. But that they do these tests. We are now like that Chinese fisher on the river shores.

Someone said the R1b must have arrived with the Proto Armenian speakers (therefore not "native" and Indo European) I clarified that by 2500 BC no known Indo European language was spoken. I showed that diversity of R1b is actually found in non Armenian speakers such as Iranic tribes Or the Northeast Caucasic speakers, to which Hurro-Urartaen has affinity (who lived in modern day Armenia for that matter).

And I also showed that the genetic make up of the Bronze Age Armenian was by 70% typical EEF. by having the West Mediterranen Atlantic_Med which Yamna lacked.

That is the whole point.

R1b in my opinion has been at least in the region between the Zagros/Albroz and Taurus region for a LONG time, if not maybe evolved there.

The Armenian language with 1200 BC is still one of the oldest in the region, thats earlier than Turkic and Semitic and slightly younger than Indo_Iranian and Hittite. And makes genetically also sense.

Since the Bronze Age Armenian was genetically closest in this order, 1. North Caucasic speakers 2. Iranic speakers 3. Armenian speakers

with a chunk of "European" DNA which in fact presents a native pre Semite Farmer ancestry

It's true that the Kyjatice culture, to which BR2 from Gamba et al. belonged, is part of the Urnfield cultures, or more precisely of the southeastern group of Urnfield cultures. However, I don't think the beginning of the Urnfield cultures is in a direct relation with the Bronze Age collapse in the Eastern Mediterranean. Because the beginning of the Urnfield cultures is now dated to the late 14th century, this is preceding the Bronze Age collapse which dates to the late 13th and early 12th century. Occasionaly, cremation followed by the burial of the ashes in urns had already occured in the pre-Lusatian culture, and it was already common in the early Lusatian culture. Both "cultures" (actually they are rather different phases of the same culture) preceded the Urnfield cultures and fall into the Middle Bronze Age. Vast urnfields are also known from the Piliny culture in northern Hungary and southern Slovakia, which also dates to the Middle Bronze Age.

Kurti North East Caucasic languages has two branch. One is Nakh who is clearly a J2a founder effect. The Hurrian link to NE Caucasian is based most importantly to the Nakh branch. So if You believe in this theory this mean that the most probable language of J2 was the Hurrian because modern day Nakhs are very high in J2a. The Daghestani group on the other side is high in J1. Avars are 65%, Lezgyns are 40% J1. Dargins have 85% of J1. So thinking that their language is related to R1b (for which only Lezgins are high, that is easily explained by their late invasions in Shirvan and Aran ) is quite difficult. That is my point.

Concerning the age of R1b in Armenia. Hamilton calculated that his TMRCA is only 3300 BC. So if he is right and I don't think he is wrong You will not find any R1b-Z2103 in Armenia before the Kura-Arax period. So the question who they are is not an easy question as one must think.